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Next: Reduction to the Membrane Up: A model for the Previous: Introduction

 

Basic Equations

Biophysical membrane excitation equations are of the form:

  eqnarray28

where u=u(t) is the transmembrane voltage, C is specific membrane capacitance, f is transmembrane current density, vector v=v(t) describes the fast gating variables, and vector w=w(t) comprises slow gating variables and intra- and extra-cellular ionic concentrations, and g and h describe their kinetics. The variables u and v have comparable characteristic times.

We begin with a bidomain approach, for a single isolated cell with an intracellular domain , tex2html_wrap_inline837 , external domain, tex2html_wrap_inline839 , and the membrane surface, tex2html_wrap_inline841 , and introduce the electrostatic potential tex2html_wrap_inline843 and tex2html_wrap_inline845 in tex2html_wrap_inline837 and tex2html_wrap_inline839 , tex2html_wrap_inline851 and tex2html_wrap_inline853 as limit values of tex2html_wrap_inline843 and tex2html_wrap_inline845 at tex2html_wrap_inline841 , and electric charge densities tex2html_wrap_inline861 and tex2html_wrap_inline863 at the inside and outside surface of the membrane.

1) Intracellularly, electroneutrality of the cytoplasm means for tex2html_wrap_inline865 ,

  equation39

for a scalar specific conductivity tex2html_wrap_inline867 .

2) In the extracellular domain, electroneutrality means for tex2html_wrap_inline869 :

  equation44

and for a homogeneous external field,

  equation47

In the case of a tissue of cells, Eq. (4) is to be replaced - e.g. for cells in a regular 3-dimensional grid, the domain tex2html_wrap_inline871 forms the elementary volume of the grid, and (4) is to be replaced by periodic boundary conditions for the ``own'' cell potential

displaymath873

or the ``oscillatory component'' of the potential, in terminology of Plonsey & Barr [1986] and Krassowska et al. [1990].

3) In the membrane, the boundary conditions of the ``volume equations'' (2,3) for tex2html_wrap_inline875 :

   eqnarray66

and surface balance of charges on the interior and exterior sides of the membrane;

   eqnarray70

here f, is the transmembrane current, tex2html_wrap_inline879 are specific conductivities, and tex2html_wrap_inline881 is the Laplacian operator on the membrane surface. For simplicity of notations, we assume here that tex2html_wrap_inline879 are constant. Electroneutrality of a membrane element gives

  equation78

The membrane capacitance is

  equation81

and transmembrane voltage is

  equation84

After defining tex2html_wrap_inline885 , through local values of u, v and w, and local kinetic equations for v and w from (1), these equation form a closed system, which determines evolution of the distribution of electric properties over the cell at given tex2html_wrap_inline897 , and so describe the action of the external electric field onto the cell.


next up previous
Next: Reduction to the Membrane Up: A model for the Previous: Introduction

Vadim Biktashev
Fri Mar 28 21:26:28 GMT 1997